Links for Palaeobotanists

Home / Introductions to both Fossil and Recent Plant Taxa / Angiosperms

Cyanobacteria and Stromatolites
Seed Plants in General
! Herbaria@
! Parasitic Plants@
! Taxonomy and Plant Classification Databases@
Plant Photographs@
Image Collections@
Picture Search@
! Trees@


A. Ali et al. (2024): A new permineralized Corypha-type coryphoid palm stem from K-Pg of India: Anatomy, systematics, saprophytic fungi, and paleoecology. Free access, Turkish Journal of Botany, 48: 105-119. doi:10.55730/1300-008X.2799.

A. Antonelli et al. (2015): An engine for global plant diversity: highest evolutionary turnover and emigration in the American tropics. In PDF, American tropics. Front. Genet., 6. doi: 10.3389/fgene.2015.00130
See also here.

! APG IV (2016): An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV. Open access, Botanical Journal of the Linnean Society, 181: 1–20.

! APG III (2009), compiled by B. Bremer et al.: An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III. In PDF, Botanical Journal of the Linnean Society, 161: 105–121. See also here.

Hank Art et al., Williams College, Biology Dept., Williamstown MA: Field botany. Go to: Evolutionary Botany. Powerpoint download and links to aricles. See especially:
Early Land Plants.
Fossil Angiosperms.
Introduction to the Angiosperms.
Powerpoint presentations.

! J. Asar et al. (2022): Early diversifications of angiosperms and their insect pollinators: were they unlinked? Free access. Trends in Plant Science, 27: 858-869. See also here.
Note figure 1: Emergence of crown angiosperms and insect pollinators.
Figure 2. Phylogeny of seed plants, depicting pollination modes of both extinct and extant lineages.

! Lorna Ash & Heather Kroening, Department of Biological Sciences, University of Alberta: Instructional Multimedia, Multimedia Topics, Botany. Snapshot taken by the Internet Archive´s Wayback Machine. Go to: Life Cycle of an Angiosperm. See also here. Online and downloadable flash 4 movies. Excellent!

B.A. Atkinson (2020): Fossil evidence for a Cretaceous rise of the mahogany family. Free access, American Journal of Botany, 107: 139-147. See also here (Science Daily), and there (PDF file, PhysOrg).

L. Augusto et al. (2014): The enigma of the rise of angiosperms: can we untie the knot? In PDF, Ecology Letters.

Francisco J. Ayala, Walter M. Fitch, and Michael T. Clegg (eds.; 2000): Variation and Evolution in Plants and Microorganisms: Toward a New Synthesis 50 Years after Stebbins. Online book, National Academy of Sciences (2000).
This expired link is now available through the Internet Archive´s Wayback Machine.
See also here

F.J. Ayala et al. (2000): Variation and evolution in plants and microorganisms: Toward a new synthesis 50 years after Stebbins. PNAS, 97: 6941-6944. Scroll to: "Trends and Patterns in Plant Evolution".

J. Baczynski and R. Claßen-Bockhoff (2023): Pseudanthia in angiosperms: a review. Free access, Annals of Botany, 132: 179–202.
"... Pseudanthia or ‘false flowers’ are multiflowered units that resemble solitary flowers in form and function.
[...] This review synthesizes historical and current concepts on the biology of pseudanthia ..."

J. Barba-Montoya et al. (2018): Constraining uncertainty in the timescale of angiosperm evolution and the veracity of a Cretaceous Terrestrial Revolution. In PDF, New Phytologist, 218: 819-834.
See also here.
Note fig. 6: The time tree of tracheophytes encompassing uncertainty of calibration strategies.
"... We reject a post-Jurassic origin of angiosperms, supporting the notion of a cryptic early history of angiosperms ..."

C.C. Baskin and J.M. Baskin (2023): The rudimentary embryo: an early angiosperm invention that contributed to their dominance over gymnosperms. Free access, Seed Science Research, 33: 63–74. S0960258523000168.
Note table 1: Information about fossil ovules, seeds and embryos of gymnosperms from the Upper Devonian to Late Cretaceous.
"... we explore the origin of the rudimentary embryo, its relationship to other kinds of plant embryos and its role in the diversification of angiosperms.
[...] We conclude that the rudimentary embryo was one of many new developments of angiosperms that contributed to their great success on earth ..."

R.M. Bateman (2020): Hunting the Snark: the flawed search for mythical Jurassic angiosperms. In PDF, Journal of Experimental Botany, 71: 22–35. See also here.

! R.M. Bateman et al. (2006): Morphological and molecular phylogenetic context of the angiosperms: contrasting the ‘top-down’ and ‘bottom-up’ approaches used to infer the likely characteristics of the first flowers. Free access, Journal of Experimental Botany, Vol. 57, No. 13, pp. 3471–3503. Major Themes in Flowering Research Special Issue.
Note fig. 1C: Male and female reproductive structures of Caytonia (Caytoniales).
Fig. 1D: Reproductive structure of Williamsoniella (Bennettitales).

J.M. Beaulieu et al. (2015): Heterogeneous rates of molecular evolution and diversification could explain the Triassic age estimate for angiosperms. Abstract.

J.M. Beaulieu et al. (2013): A Southern Hemisphere origin for campanulid angiosperms, with traces of the break-up of Gondwana. In PDF, BMC Evolutionary Biology, 13.

C.M. Belcher and V.A. Hudspith (2017): Changes to Cretaceous surface fire behaviour influenced the spread of the early angiosperms. New Phytologist, 213: 1521–1532.

! C.D. Bell et al. (2010): The age and diversification of the angiosperms re-revisited. Free access, American Journal of Botany, 97: 1296-1303.

! M.J. Benton et al. (2022): The Angiosperm Terrestrial Revolution and the origins of modern biodiversity. Free access, New Phytologist, 233: 2017–2035.
Note fig. 1: Evolution of hyperdiverse terrestrial life.
Fig. 3: Key stages in Earth history and angiosperm evolution through the Angiosperm Terrestrial Revolution.
Also worth checking out:
Flowering plants: an evolution revolution. (Univ. of Bristol, November 17, 2021).
How 'Flower Power' Quite Literally Transformed Earth Millions of Years Ago (by T. Koumoundouros, January 08,2022).

Museum of Paleontology, University of California (UCMP), Berkeley, CA: Introduction to the Anthophyta, Monocots versus Dicots, and Introduction to the Liliopsida. The Monocots.

! Museum of Paleontology, University of California, Berkeley:
The Cleared Leaf Collection. Excellent!
An image gallery of modern leaves that have been bleached and stained to make their venation patterns more visible. Leaf shape, venation, and features of the margin, base and apex constitute important taxonomic and physiognomic characters.
You can search the collection from the
Paleontology Collections Photos page or
the collection at the Modern Cleared Leaf Photos page.
Don't miss the helpful
! Manual of Leaf Architecture. In PDF.

S. Beurel et al (2024): First flower inclusion and fossil evidence of Cryptocarya (Laurales, Lauraceae) from Miocene amber of Zhangpu (China). In PDF, Fossil Record, 27: 1–11.
See likewise here and there.
"... We here described the first Cenozoic Lauraceae flower of Asia and confirmed the presence of Cryptocarya in the Miocene Zhangpu flora
[...] We scanned the specimen using synchrotron radiation-based micro-computed tomography (SRìCT) and then compared the fossil with extant flowers of the genus ..."

S. Bhadra et al. (2023): From genome size to trait evolution during angiosperm radiation. In PDF, Trends in Genetics.
See likewise here.
Note figure 1: Schematic representation illustrating the relationship between genome size change and trait evolution.

H.J.B. Birks (2020): Angiosperms versus gymnosperms in the Cretaceous. Open access, PNAS, 117: 30879-30881.

J. Blanchard et al. (2016): Fruits, seeds and flowers from the Bovay and Bolden clay pits (early Eocene Tallahatta Formation, Claiborne Group), northern Mississippi, USA. In PDF, Palaeontologia Electronica. See also here.

W.J. Bond and A.C. Scott (2010): Fire and the spread of flowering plants in the Cretaceous. In PDF, New Phytologist, 188: 1137-1150.

! H. Boukhamsin et al. (2023): Early Cretaceous angiosperm radiation in northeastern Gondwana: Insights from island biogeography theory. Free access, Earth-Science Reviews, 242.

C.K. Boyce and A.B. Leslie (2012): The Paleontological Context of Angiosperm Vegetative Evolution. In PDF, International Journal of Plant Sciences, 173: 561–568.
See also here.
"... a survey of the fossil record demonstrates that most anatomical traits that are now unique to the angiosperms were more broadly distributed among extinct lineages.
[...] Of all the various vegetative morphological traits that have been traditionally linked to angiosperm success, only the evolution of very high leaf vein densities appears to be truly unique to the angiosperms. ..."

C.K. Boyce et al. (2010): Angiosperms Helped Put the Rain in the Rainforests: The Impact of Plant Physiological Evolution on Tropical Biodiversity. PDF file, Annals of the Missouri Botanical Garden, 97: 527-540.
Provided by the Internet Archive´s Wayback Machine.

C.K. Boyce et al. (2009). Angiosperm leaf vein evolution was physiologically and environmentally transformative. PDF file, Proceedings of the Royal Society B, 276: 1771-1776.

Jamie Boyer, The New York Botanical Garden: Welcome to the Botanical Education site. Here you find information on courses and blogs created by Jamie Boyer, e.g.
! Plant Diversity. About the diversity, morphologies, life histories, and evolution of plants, as well as information about bacteria, archaea, fungi, and algae/protists.

J.D. Boyko et al. (2023): The evolutionary responses of life-history strategies to climatic variability in flowering plants. Free access, New Phytologist, doi: 10.1111/nph.18971.
See also here (in PDF).
Note figure 1: Global distribution of vascular plant diversity and proportion of annual plants.

J. Bres et al. (2021): The Cretaceous physiological adaptation of angiosperms to a declining pCO2: a modeling approach emulating paleo-traits. Free access, Biogeosciences, 18: 5729–5750.
"... we show that protoangiosperm physiology does not allow vegetation to grow under low pCO2
[...] confirms the hypothesis of a likely evolution of angiosperms from a state of low leaf hydraulic and photosynthetic capacities at high pCO2 to a state of high leaf hydraulic and photosynthetic capacities linked to leaves with more and more veins together ..."

The palaeofiles. Articles here have all been prepared by students on the palaeobiology programmes in Bristol:
! Purported Triassic angiosperms.
Now provided by the Internet Archive´s Wayback Machine.

The palaeofiles. Articles here have all been prepared by students on the palaeobiology programmes in Bristol:
! The origin and evolution of angiosperms.
Now provided by the Internet Archive´s Wayback Machine.

T.J. Brodribb and T.S. Feild (2010): Leaf hydraulic evolution led a surge in leaf photosynthetic capacity during early angiosperm diversification. In PDF, Ecology Letters, 13: 175-183. See also here.
"... Our data suggest that early terrestrial angiosperms produced leaves with low photosynthetic rates, but that subsequent angiosperm success is linked to a surge in photosynthetic capacity during their early diversification".

G.E. Budd et al. (2021): Fossil data do not support a long pre-Cretaceous history of flowering plants. Free access, bioRxiv.

G.E. Budd et al. (2021): The use of geological and paleontological evidence in evaluating plant phylogeographic hypotheses in the Northern Hemisphere Tertiary. Free access, See also here (in PDF).

Benjamin Burger, Utah State University, Vernal, Utah:
Why study fossil plants?
Invertebrate Paleontology and Paleobotany.
How did plants colonize the land, based on the fossil record?
How did the first seed plants (the Gymnosperms) evolve?
How did gymnosperms diversify during the early Mesozoic to become a modern dominate plant group?
How good is the fossil record of Cycads?
What is the significance of the fossil record of Ginkgo?
What is the fossil record of Horsetails?
Fossil Algae.
! What is an Angiosperm?
Video lectures.

! R.J. Burnham (2009): An overview of the fossil record of climbers: bejucos, sogas, trepadoras, lianas, cipós, and vines. PDF file, Rev. bras. paleontol., 12: 149-160.
Snapshot provided by the Internet Archive´s Wayback Machine.

R.J. Burnham (1994): Patterns in tropical leaf litter and implications for angiosperm paleobotany. In PDF, Review of Palaeobotany and Palynology.

R.J. Butler et al. (2009): Diversity patterns amongst herbivorous dinosaurs and plants during the Cretaceous: implications for hypotheses of dinosaur/angiosperm co-evolution. Free access, Journal of Evolutionary Biol., 22: 446-459.

! A.S. Chanderbali et al. (2016): Evolving Ideas on the Origin and Evolution of Flowers: New Perspectives in the Genomic Era. In PDF, Genetics, 202: 1255–1265. See also here.

G. Chomicki et al. (2017): Evolution and ecology of plant architecture: integrating insights from the fossil record, extant morphology, developmental genetics and phylogenies. Annals of Botany 120: 855–891. See also here (in PDF).
Note Fig. 9: A timeline of plant architectures and branching mechanisms through time.

! Michael Clayton, Department of Botany, University of Wisconsin, Madison: Instructional Technology (BotIT). Some image collections. Excellent! Go to:

C. Coiffard et al. (2023): The emergence of the tropical rainforest biome in the Cretaceous. Free access, Biogeosciences, 20: 1145–1154.
See also: Einem frühen Regenwald auf der Spur. In German,

C. Coiffard et al. (2012): Rise to dominance of angiosperm pioneers in European Cretaceous environments , Abstract. See also here ( and there (

C. Coiffard et al. (2012): Deciphering Early Angiosperm Landscape Ecology Using a Clustering Method on Cretaceous Plant Assemblages. In PDF.

M. Coiro et al. (2024): Parallel evolution of angiosperm-like venation in Peltaspermales: a reinvestigation of Furcula. Open access, New Phytologist, doi: 10.1111/nph.19726.
"... Although a hierarchical-reticulate venation also occurs in some groups of extinct seed plants, it is unclear whether these are stem relatives of angiosperms
[...] We further suggest that the evolution of hierarchical venation systems in the early Permian, the Late Triassic, and the Early Cretaceous represent ‘natural experiments’ that might help resolve the selective pressures enabling this trait to evolve ..."

! M. Coiro et al. (2019): How deep is the conflict between molecular and fossil evidence on the age of angiosperms? Free access, New Phytologist, doi: 10.1111/nph.15708.
"... Critical scrutiny shows that supposed pre-Cretaceous angiosperms either represent other plant groups or lack features that might confidently assign them to the angiosperms. ..."

! F.L. Condamine et al. (2020): The rise of angiosperms pushed conifers to decline during global cooling. Free access, Proceedings of the National Academy of Sciences, 117: 28867–28875.
Note figure 1: An overview of hypothetical determinants of conifer diversification over time.
Figure 2: Global diversification of conifers inferred from a molecular phylogeny and the fossil record.
Figure 3: Drivers of conifer diversification dynamics.

B. Cornet (1989): The reproductive morphology and biology of Sanmiguelia lewisii, and its bearing on angiosperm evolution in the Late Triassic. Evolutionary trends in Plants.

B. Cornet: Why do Paleobotanists Believe in a Cretaceous Origin of Angiosperms? A controversial topic. This website presents palaeobotanical evidence on the origin of flowering plants, with evidence for and against a Cretaceous origin. See also: Angiosperm Evolution.
Websites still available via Internet Archive Wayback Machine.

! B. Crair (2023): The Fossil Flowers That Rewrote the History of Life. Free access, The New Yorker.
"... Instead of breaking rocks, she crumbled soft sediments into a sieve, washed away the sand grains in water, and saved the tiny specks of charcoal that were left behind.
[...] Fresh discoveries, she added, could radically change the known history of flowers.
[...] “A day in the field can be years of work in the laboratory.” ..."

! P.R. Crane and A.B. Leslie (2013): Major Events in the Evolution of Land Plants. In PDF. The Princeton Guide to Evolution.
1. Phylogenetic framework.
2. Origin and diversification of land plants.
3. Origin and diversification of vascular plants.
4. Origin and diversification of seed plants.
5. Origin and diversification of flowering plants.
6. Innovation in the land plant body.
7. Innovation in land plant reproduction.
8. Co-evolution with animals.
9. Patterns of extinction.
See also here, and there (Google books).

P.R. Crane et al. (2010): Darwin and the Evolution of Flowers. PDF file, Phil. Trans. R. Soc. B, 365: 347-350.

! W.L. Crepet and K.J. Niklas (2009): Darwin´s second "abominable mystery": Why are there so many angiosperm species? Open access, American Journal of Botany, 96: 366-381.

W.L. Crepet (2008): The Fossil Record of Angiosperms: Requiem or Renaissance? Abstract, Annals of the Missouri Botanical Garden, 95: 3-33.
See also here.
"... a reasonably good fossil record of angiosperms is emerging
[...] One of its most striking aspects is the rapid radiation of angiosperm taxa that are now unusually diverse around two particular times in geological history: the Turonian and Early Tertiary. Possible reasons for these intervals of rapid radiation among angiosperms will be discussed.


! W.L. Crepet et al. (2004): Fossil evidence and phylogeny: the age of major angiosperm clades based on mesofossil and macrofossil evidence from Cretaceous deposits. Free access, American Journal of Botany, 91: 1666-1682.
! Beautifully preserved charcoalified flowers!

William L. Crepet, Department of Plant Biology, Cornell University, Ithaca, NY: Progress in understanding angiosperm history, success, and relationships: Darwin's abominably "perplexing phenomenon". PNAS 2000; 97: 12939-12941.

Judith L. Croxdale, Department of Botany, University of Wisconsin, Madison (website hosted by Biology Online): Stomatal patterning in angiosperms. Stomatal pattern types, means of measuring them, advantages of each type of measurement, and then present patterning from evolutionary, physiological, ecological, and organ views are discussed.
Website outdated, a version archived by the Internet Archive Wayback Machine.

D.-F. Cui et al. (2022): A Jurassic flower bud from China. In PDF, Geological Society, London. See also here.
"... The leaf scars are 0.4–0.6 mm wide and 0.23 mm thick, with terminal abscission zones (Fig. 3b, d & e). ..."

Charles Daghlian (Dartmouth College, Hannover, NH) and Jennifer Svitko, Paleobotanical Holdings at the Liberty Hyde Bailey Hortorium at Cornell University: Paleoclusia 3D Reconstructions. Movies from CT scans done on the Turonian fossils. See also here (W.L. Crepet and K.C. Nixon 1998, abstract and photos).
Snapshots provided by the Internet Archive´s Wayback Machine.

M.J. Dallwitz Australia (DELTA – DEscription Language for TAxonomy).
Descriptions, illustrations, interactive identification, and information retrieval from DELTA databases.

H.J. de Boer et al. (2012): A critical transition in leaf evolution facilitated the Cretaceous angiosperm revolution. In PDF, Nature Communications, 3. See also here.

! D.L. Dilcher (2001): Paleobotany: some aspects of non-flowering and flowering plant evolution. In PDF, Taxon.
See also here.
For early angiospermous fossil floras see figure 1 (on PDF page 5).

David Dilcher (2000): Toward a new synthesis: Major evolutionary trends in the angiosperm fossil record. PDF file, Proc Natl Acad Sci U S A., 97: 7030-7036. See also here.

! D.L. Dilcher (1974): Approaches to the identification of angiosperm leaf remains. In PDF, The Botanical Review, 40: 1–157.
See also here.
"... Many techniques for the study of the morphology of modern and fossil leaves are included in this paper as well as tables outlining features of leaf venation and the epidermis ..."

D. Dimitrov et al. (2023): Diversification of flowering plants in space and time. Free access, Nature Communications, 14.
"... Using a newly generated genus-level phylogeny and global distribution data for 14,244 flowering plant genera, we describe the diversification dynamics of angiosperms through space and time. Our analyses show that diversification rates increased throughout the early Cretaceous and then slightly decreased or remained mostly stable until the end of the Cretaceous–Paleogene mass extinction event 66 million years ago. After that, diversification rates increased again towards the present ..."

P. Donoghue (2019): Evolution: The Flowering of Land Plant Evolution. Abstract, Current Biology. See also:
Evolution: The flowering of land plant evolution - whence and whither? (in PDF).

M.J. Donoghue and J.A. Doyle (2000): Seed plant phylogeny: Demise of the anthophyte hypothesis?. Free access, Current Biology, 10: R106-R109.
See also here.
"... Recent molecular phylogenetic studies indicate, surprisingly, that Gnetales are related to conifers, or even derived from them ..."

Stephen R. Downie and Kenneth R. Robertson, Life Sciences at the University of Illinois, Urbana-Champagne: Systematics of Plants. This course introduces the principles and methods of identifying, naming, and classifying flowering plants. It includes a survey of selected flowering plant families and provides information on their interrelationships. Go to: Digital Flowers.
These expired links are available through the Internet Archive´s Wayback Machine.

J.A. Doyle and P.K. Endress (2014): Integrating Early Cretaceous Fossils into the Phylogeny of Living Angiosperms: ANITA Lines and Relatives of Chloranthaceae Int. J. Plant Sci., 175: 555–600. See also here.

! J.A. Doyle (2012): Molecular and fossil evidence on the origin of angiosperms. In PDF, Annual Review of Earth and Planetary Sciences, 40: 301-26.

! J.A. Doyle (1998): Molecules, morphology, fossils, and the relationship of angiosperms and Gnetales. In PDF, Molecular phylogenetics and evolution.

X.Y. Du et al. (2021): Simultaneous diversification of Polypodiales and angiosperms in the Mesozoic. In PDF, Cladistics, 37.
See also here.
Note fig. 2: Summary chronograms of Polypodiales.
Fig. 5. Comparison of lineage through time plots for Polypodiales and angiosperms.
"... The estimated divergence patterns of Polypodiales and angiosperms converge to a scenario in which their main lineages were established simultaneously shortly before the onset of the Cretaceous Terrestrial Revolution ..."

M. Eberlein (2015): Bestimmungs- und Verbreitungsatlas der Tertiärflora Sachsens – Angiospermenblätter und Ginkgo. PDF file (in German). Thesis, University of Dresden (in German). First part of a reference book of the Tertiary flora of Saxony.
See also here.

! P.K. Endress (2011): Angiosperm ovules: diversity, development, evolution. Free access, Annals of Botany, 107: 1465-1489.

P.K. Endress and J.A. Doyle (2009): Reconstructing the ancestral angiosperm flower and its initial specializations. Free access, American Journal of Botany, 96: 22-66.

! Mike Farabee, Estrella Mountain Community College Center, Avondale, Arizona:
On-Line Biology Book. Introductory biology lecture notes. Go to:
Flower Structure.
Fertilization and Fruits.
These expired links are available through the Internet Archive´s Wayback Machine.

J.A. Fawcett and Y. Van de Peer (2010): Angiosperm polyploids and their road to evolutionary success. Trends in Evolutionary Biology.
See also here.

T.S. Feild et al. (2011): Fossil evidence for Cretaceous escalation in angiosperm leaf vein evolution. In PDF, PNAS, 108: 8363-8366.

Taylor S. Feild and N.C. Arens (2007): The ecophysiology of early angiosperms. PDF file, Plant, Cell and Environment, 30: 291-309.

! T.S. Feild et al. (2004): Dark and disturbed: a new image of early angiosperm ecology. PDF file, Paleobiology, 30: 82-107.
See also here.
"... The vegetative flexibility that evolved in the understory, however, may have been a key factor in their diversification in other habitats. Our inferences based on living plants are consistent with many aspects of the Early Cretaceous fossil record ..."

C. Ferrándiz et al. (2010): Carpel development. In PDF, Advances in Botanical Research, 55: 1-73.
See also here.

S. Fields (2021): Diversification of Angiosperms During the Cretaceous Period. In PDF, Undergraduate Student Theses, Environmental Studies Program at DigitalCommons@University of Nebraska,Lincoln. See also here.

! C.S.P. Foster (2016): The evolutionary history of flowering plants. In PDF, Journal & Proceedings of the Royal Society of New South Wales, 149: 65-82.

William Friedman et al., Department of Ecology and Evolutionary Biology, University of Colorado, Boulder: Molecular and Organismal Research in Plant History, MORPH. MORPH, an NSF research coordination network, fosters cross-disciplinary interactions between organismic and molecular plant biologists studying the evolution of morphological diversity to promote a modern synthesis in plant evolutionary developmental biology. See also: Publications.
A version archived by the Internet Archive´s Wayback Machine.

! E.M. Friis et al. (2019): The Early Cretaceous Mesofossil Flora of Torres Vedras (Ne of Forte Da Forca), Portugal: A Palaeofloristic Analysis of an Early Angiosperm Community. Open access, Fossil Imprint, 75: 153–257. See also here (in PDF).
"... the oldest mesofossil flora containing angiosperm remains to be described in detail based on well-preserved flower, fruit and seed remains. It provides the most detailed information currently available on the structural diversity of angiosperms at this early stage in their evolution, the range of angiosperm species present, and their relationships to extant angiosperm lineages. ..."

E.M. Friis et al. (2015): Exceptional preservation of tiny embryos documents seed dormancy in early angiosperms. In PDF, Nature, 528: 551-554. See also here.

! E.M. Friis et al. (2011): Early Flowers and Angiosperm Evolution. Abstract, Cambridge University Press.
See also here (in PDF, long download time) and there (Google books).
Also worth checking out: Book Review, by P.J. Rudall, Botanical Journal of the Linnean Society, 170. In PDF.
"... This long-awaited book represents not only a remarkable tour de force of palaeobotanical literature, but also a potentially enduring biological textbook. ..."

Else Marie Friis, Swedish Museum of Natural History, Stockholm: Cretaceous angiosperms from Europe and North America (Silvianthemum suecicum), and Cretaceous angiosperms from Kazakhstan.
Snapshots taken by the Internet Archive´s Wayback Machine.

! Else Marie Friis et al. (2010): Diversity in obscurity: fossil flowers and the early history of angiosperms. PDF file, Phil. Trans. R. Soc. B, 365: 369-382. Some of the specimens are charcoalified and have retained their original three-dimensional shape. See also here.

E.M. Friis et al. (2006): Cretaceous angiosperm flowers: Innovation and evolution in plant reproduction. Abstract, Palaeogeography, Palaeoclimatology, Palaeoecology, 232: 251-293.

! M.W. Frohlich & M.W. Chase (2007): After a dozen years of progress the origin of angiosperms is still a great mystery. In PDF, Nature, 450: 1184-1189.
See also here.

! Q. Fu et al. (2023): Micro-CT results exhibit ovules enclosed in the ovaries of Nanjinganthus. Open access, Scientific Reports, 13.
Note figure 4: Micro-CT results exhibit ovules enclosed in the ovaries of Nanjinganthus.

Q. Fu et al. (2018): An unexpected noncarpellate epigynous flower from the Jurassic of China. In PDF, eLife, 7: e38827. See also:
D.W. Taylor and H. Li (2019): Paleobotany: Did flowering plants exist in the Jurassic period.

Q. Fu et al. (2017): Nanjinganthus: An Unexpected Flower from the Jurassic of China. In PDF, bioRxiv (pronounced "bio-archive"). See also here.

J.F. Genise et al. (2020): 100 Ma sweat bee nests: Early and rapid co-diversification of crown bees and flowering plants. Open access, PLoS ONE 15: e0227789.

N. Gentis et al. (2024): First fossil woods and palm stems from the mid-Paleocene of Myanmar and implications for biogeography and wood anatomy. Open access, Am J Bot., 111.

W.V. Gobo et al. (2023): A new remarkable Early Cretaceous nelumbonaceous fossil bridges the gap between herbaceous aquatic and woody protealeans. Open access, Scientific Reports, 13.
Note figure 9: Reconstruction of Notocyamus hydrophobus gen. nov. et sp. nov. in its likely environment.

W.V. Gobo et al. (2022): First evidence of ranunculids in Early Cretaceous tropics. Open access, Scientific Reports, 12.

B. Gomez et al. (2015): Montsechia, an ancient aquatic angiosperm. In PDF, PNAS, 112: 10985–10988. See alao here.
Note Fig. 3: Reconstructions of Montsechia vidalii.

! R. Gorelick (2001): Did insect pollination cause increased seed plant diversity? PDF file, Biological Journal of the Linnean Society, 74: 407-427.

J. Gravendyck et al. (2022): Early Angiosperms - How far can we reliably go back in the pollen record. Abstract, 11th European Palaeobotany and Palynology Conference Abstracts, Program and Proceedings, Swedish Museum of Natural History, Stockholm.

! G. Han et al. (2016): A Whole Plant Herbaceous Angiosperm from the Middle Jurassic of China. In PDF, Acta Geologica Sinica. See also here (abstract) and there (in German, with photograph and reconstruction).

Ben Harder, National Geographic News: Dino-Era Fossil—The First Flower? About the Archaefructaceae.

! P.S. Herendeen et al. (2017): Palaeobotanical redux: revisiting the age of the angiosperms. In PDF, Nature Plants 3. See also here.

L. Hernandez-Sandoval et al. (2023): Nichima gen. nov. (Alismataceae) based on reproductive structures from the Oligocene]Miocene of Mexico. Open access, American Journal of Botany, 110.
Note figure 7: Fossil flower reconstructions and proposed inflorescence organization.
"... Two fossil flowers preserved in amber from the Miocene ..."

T.E. Higham et al. (2022): The Evolution of Mechanical Properties of Conifer and Angiosperm Woods. In PDF, Integrative and Comparative Biology, 62: 668–682.
See also here.

P.A. Hochuli and S. Feist-Burkhardt (2013): Angiosperm-like pollen and Afropollis from the Middle Triassic (Anisian) of the Germanic Basin (Northern Switzerland). In PDF, Frontiers in plant science.

P.A. Hochuli, Paläontologisches Institut und Museum, Universität Zürich: Timing of angiosperm evolution. Research project description.

Natalia Holden, Department of Biological Sciences, University of Alberta, Edmonton, Canada: The early Angiosperms: Paleophytogeography and Depositional Settings. A slideshow.
This expired link is available through the Internet Archive´s Wayback Machine.

! S. Hu et al. (2008): Early steps of angiosperm-pollinator coevolution. PDF file, PNAS, 105: 40-245. See also here (abstract).

I.B. Huegele and S.R. Manchester (2020): An Early Paleocene Carpoflora from the Denver Basin of Colorado, USA, and Its Implications for Plant-Animal Interactions and Fruit Size Evolution. Free access, Int. J. Plant Sci., 181: 646–665.

! Norman F. Hughes (1994): The Enigma of Angiosperm Origins. 405 pages. Provided by Cambridge University Press through the Google Print Publisher Program.
See also here.

Norman F. Hughes (1982): Palaeobiology of Angiosperm Origins: Problems of Mesozoic Seed-Plant Evolution. Provided by Google books.

Marty Huss: Angiosperm Life Cycle. Powerpoint presentation.

N.A. Jud et al. (2018): A new fossil assemblage shows that large angiosperm trees grew in North America by the Turonian (Late Cretaceous). In PDF, Sci. Adv., 4: eaar8568.
"A large silicified log (maximum preserved diameter, 1.8 m; estimated height, ca. 50 m) is assigned to the genus Paraphyllanthoxylon; it is the largest known pre-Campanian angiosperm and the earliest documented occurrence of an angiosperm tree more than 1.0 m in diameter."

N.A. Jud (2015): Fossil evidence for a herbaceous diversification of early eudicot angiosperms during the Early Cretaceous. In PDF, Proc. R. Soc., B, 282. See also here.

N.A. Jud and L.J. Hickey (2013): Potomacapnos apeleutheron gen. et sp. nov., a new Early Cretaceous angiosperm from the Potomac Group and its implications for the evolution of eudicot leaf architecture. In PDF, Am. J. Bot., see also here.

! O. Katz (2018): Extending the scope of Darwin’s ‘abominable mystery’: integrative approaches to understanding angiosperm origins and species richness. Open access, Annals of Botany, 121: 1–8.
See also here (Botany One).

A.J. Kerkhoff et al. (2014): The latitudinal species richness gradient in New World woody angiosperms is consistent with the tropical conservatism hypothesis. In PDF, PNAS, 111: 8125–8130. See also here.

! Kimball´s Biology Pages (by John W. Kimball). The pages represent an online biology textbook. Go to: Sexual Reproduction in Angiosperms.

Michael Knee, Department of Horticulture and Crop Science, Ohio State University:
General Plant Biology, Horticulture and Crop Science 300, Online Resources. Go to: ANTHOPHYTA I,
Evolution of flowering plants, and

! Knox College, Galesburg, Illinois: The Seed Plants: Gymnosperms & Angiosperms. Lecture note, Powerpoint presentation.
This expired link is still available through the Internet Archive´s Wayback Machine.

Ari and Susan Kornfeld, Natural Perspective: The Plant Kingdom: Dicots Overview. A version archived by Internet Archive Wayback Machine.

V. Krassilov and S. Barinova (2014): "Flower" of Magnolia grandiflora is not flower and what about "basal angiosperms". In PDF, Journal of Plant Sciences, 2: 282-293.

V. Krassilov (2012): Fossil record of angiosperm origin: new evidence and interpretation. In PDF, Horizons in Earth Science Research, 8. (Nova Publishers, New York).

V.A. Krassilov (2009): Diversity of Mesozoic Gnetophytes and the First Angiosperms. PDF file, Paleontological Journal, 43: 1272-1280. A version archived by Internet Archive Wayback Machine.

A. Kremer and A.L. Hipp (2019): Oaks: an evolutionary success story. Free access, New Phytologist, doi: 10.1111/nph.16274.

B.B. Lamont and T. He (2012): Fire-adapted Gondwanan Angiosperm floras evolved in the Cretaceous. In PDF, BMC Evolutionary Biology, 12. See also here.

Gerhard Leubner Lab, University Freiburg, Germany: Seed Evolution. Go to: Angiosperm seed evolution and species diversification.

Gerhard Leubner, The Seed Biology Place, Molecular Plant Sciences, University Freiburg, Germany: Seed evolution. Origin and evolution of the seed habit. See also: Seed dictionary English-German.

Z.-J. Liu and X. Wang (2017): Yuhania: a unique angiosperm from the Middle Jurassic of Inner Mongolia, China. Open access, Historical Biology, 29: 431-441.

! H.T. Li et al. (2019): Origin of angiosperms and the puzzle of the Jurassic gap. Abstract, Nature Plants, 5: 461–470. See also here (in PDF).
"... With a well-resolved plastid tree and 62?fossil calibrations, we dated the origin of the crown angiosperms to the Upper Triassic, with major angiosperm radiations occurring in the Jurassic and Lower Cretaceous. This estimated crown age is substantially earlier than that of unequivocal angiosperm fossils, and the difference is here termed the ‘Jurassic angiosperm gap’. ..."

Biological Sciences, Ohio State University, Lima: Plant Biology at OSU Lima.
This expired link is now available through the Internet Archive´s Wayback Machine.

A. Linkies et al. (2010): The evolution of seeds. PDF file, New Phytologist.

S.A. Little et al. (2014): Reinvestigation of Leaf Rank, an Underappreciated Component of Leo Hickey´s Legacy. In PDF.

Z.J. Liu et al. (2019): Zhangwuia: an enigmatic organ with a bennettitalean appearance and enclosed ovules. In PDF, Earth and Environmental Science Transactions of The Royal Society of Edinburgh, 108 (Agora Paleobotanica), : 419-428. See also here.

Z.-J. Liu et al. (2018): The Core Eudicot Boom Registered in Myanmar Amber. Open access, Scientific Reportsvolume 8.
Note figure 5: Reconstruction of Lijinganthus revoluta.

Z.-J. Liu and X. Wang (2016): A perfect flower from the Jurassic of China. In PDF, Historical Biology, 28: 707-719. See also here (Abstract).

W.-Z. Liu et al. (2014): From leaf and branch into a flower: Magnolia tells the story. Open access, Botanical Studies, 55.

! A.M. López-Martínez et al. (2023): Angiosperm flowers reached their highest morphological diversity early in their evolutionary history. Open access, New Phytologist, 241: 1348–1360. doi: 10.1111/nph.19389.
"... Based on a comprehensive dataset focusing on 30 characters describing floral structure across angiosperms, we used 1201 extant and 121 fossil flowers to measure floral disparity and explore patterns of floral evolution through time and across lineages ..."

P.S. Soltis and D.E. Soltis (2004): The origin and diversification of angiosperms. Free access, American Journal of Botany, 91: 1614-1626.

Department of Botany, University of Wisconsin, Madison:
Plant Systematics Collection. This web site provides structured access to a teaching collection of plant images representing over 250 families and 1000 genera of vascular plants. Go to:
Phylum Magnoliophyta (Flowering Plants).
Still available via Internet Archive Wayback Machine.

S. Magallón et al. (2015): A metacalibrated time-tree documents the early rise of flowering plant phylogenetic diversity. In PDF, New Phytologist.

! S. Magallón (2009): Flowering plants (Magnoliophyta). PDF file, In: S.B. Hedges and S. Kumar (eds.): The Timetree of Life (see here).

S. Magallón and A. Castillo (2009): Angiosperm diversification through time. Free access, American Journal of Botany, 96: 349-365.

Steven R. Manchester (website hosted by International Organisation of Palaeobotany): Living Fossils, Davidia - the Dove Tree and its fossil record.
The link is to a version archived by the Internet Archive´s Wayback Machine.

J. Marugán-Lobón et al. (2022): The Las Hoyas Lagerstätte: a palaeontological look to an Early Cretaceous wetland. Open access, Journal of the Geological Society.
See also here (in PDF).
"... The site has yielded a particularly diverse assemblage of more than twenty thousand plant and animal fossils, many of which present unprecedented soft-tissue preservation, including microstructural details. Among the most significant discoveries are the oldest angiosperms, ..."

! J. Massoni et al. (2014): Fossil calibration of Magnoliidae, an ancient lineage of angiosperms. Palaeontologia Electronica.

Duane D. McKenna et al. (2009): Temporal lags and overlap in the diversification of weevils and flowering plants. PDF file, PNAS, 106: 7083-7088. See also here (abstract).

Department of Biological Sciences, University of Memphis: History of Angiosperm Classification (Post Darwin´s Theory of Evolution). In PDF.

John M. Miller (2006), School of Pure and Applied Sciences, University of the South Pacific (USP):
! Origin of Angiosperms. Go to: Insect-Plant Mutualisms.
Still available via Internet Archive Wayback Machine.

B.A.R. Mohr and H. Eklund&xnbsp;(2003): Araripia florifera, a magnoliid angiosperm from the Lower Cretaceous Crato Formation (Brazil). In PDF, Review of Palaeobotany and Palynology, 126: 279-292. See also here.
Note figure 3: Araripia florifera nov. gen. nov. spec., tentative reconstruction.

! Sebastian Molnar, Department of Zoology, University of British Columbia, Vancouver: Evolution and the Origins of Life. A directory of introductions concerning evolution, with a bias to Plant Biology and Evolution. Go to: Angiosperm Origins and Evolution.

! C. Müller et al. (2023): An integrated leaf trait analysis of two Paleogene leaf floras. In PDF, PeerJ 11: e15140
See also here.
Note figure 1: Schematic overview of the datasets used and their selection process.
Figure 6: Herbivory metrics compared between Seifhennersdorf and Suletice-Berand regarding whole assemblages and fossil-species phenology.
"... This study presents the Integrated Leaf Trait Analysis (ILTA), a workflow for the combined application of methodologies in leaf trait and insect herbivory analyses on fossil dicot leaf assemblages ..."

Nature Science Update (December 8, 1999): One for the Vine. "A prickly climbing vine", Vasovinea tianii (Gigantopteridales), that lived more than 250 million years ago could shed light on the origin of flowering plants.
This expired link is available through the Internet Archive´s Wayback Machine.

! D. Naware et al. (2024): Patterns of variation in fleshy diaspore size and abundance from Late Triassic–Oligocene. Open access, Biological Reviews, 99: 430-457.
Note figure 2: Genus richness of seed plants with fleshy and non-fleshy diaspores from Late Triassic to Oligocene across mid- to high latitudes and low latitudes.
"... Vertebrate-mediated seed dispersal is a common attribute of many living plants, and variation in the size and abundance of fleshy diaspores is influenced by regional climate and by the nature of vertebrate seed dispersers among present-day floras.
[...] We present a new data set of more than 800 georeferenced fossil diaspore occurrences spanning the Triassic–Oligocene, across low to mid- to high palaeolatitudes ..."

D.L. Nickrent (2020): Parasitic angiosperms: How often and how many? Free access, Taxon.

A.B. Nicotra et al. (2011): The evolution and functional significance of leaf shape in the angiosperms. In PDF, Functional Plant Biology, 38: 535-552. See also here.

NOVA (science series on television): First Flower.

D. Oakley et al. (2009): Morphometric analysis of some Cretaceous angiosperm woods and their extant structural and phylogenetic analogues: Implications for systematics. PDF file, Review of Palaeobotany and Palynology, 157: 375-390.
See also here.

J. Ollerton and E. Coulthard (2009): Evolution of Animal Pollination. In PDF, Science, 326.

! D. Peris and F.L. Condamine (2024): The angiosperm radiation played a dual role in the diversification of insects and insect pollinators. Open access, Nature Communications, 15.
Note figure 2: Accumulated diversification of insect families through geological time.
Figure 3: Correlation trends of different analysed drivers for origination (in blue) and extinction (in red) rates on insect diversity for two periods of time: the Angiosperm Terrestrial Revolution timeframe (100–50Ma), and for the Angiosperm Dominance period (50–0Ma).
"... Macroevolutionary studies of insect and plant diversities support the hypothesis that angiosperms diversified after a peak in insect diversity in the Early Cretaceous. Here, we used the family-level fossil record of insects as a whole, and insect pollinator families in particular, to estimate diversification rates and the role of angiosperms on insect macroevolutionary history ..."

! D. Peris and F.L. Condamine (2023): The dual role of the angiosperm radiation on insect diversification. Free access, bioRxiv.
See also here.
"... We found that, among the six tested variables, angiosperms had a dual role that has changed through time with an attenuation of insect extinction in the Cretaceous and a driver of insect origination in the Cenozoic. ..."

! D. Peris et al. (2017): False Blister Beetles and the Expansion of Gymnosperm-Insect Pollination Modes before Angiosperm Dominance. In PDF, Current Biology, 27. See also here.

! M. Philippe et al. (2008): Woody or not woody? Evidence for early angiosperm habit from the Early Cretaceous fossil wood record of Europe. PDF file, Palaeoworld, 17: 142-152.
See also here.

Ray Phillips, Information Technology Services, Colby College, Waterville, Maine: World Wide Flowering Plant Family Identification. Select the characters that are present in the specimen being identified and press "Submit". Database is part of "Biology 211: Flowering Plant Taxonomy", an introduction to the principles and practice of flowering plant taxonomy.

! Phyto Images
Database hosted by the Cornell University Vascular Plant Herbarium. Software deveopment by Kevin C. Nixon and Jan De Laet).
This site includes a wide variety of vascular plant and bryophyte photos of high quality. Phyto Images belongs to DOL (, which is a web interface based on the Encino Software Project. The Encino project is a unified set of software tools for storing, retrieving, and analyzing biodiversity. Superbly done!

! (by Max Antheunisse and Jan Koeman). is a completely non-commercial website. On top you see 2 search boxes at the right. The white one is for entering scientific names, the grey one for vernacular ones.
You may likewise navigate from:
the List of currently included artists.
Don't miss the useful link list

Tõnu Ploompuu, Biology, Tallinn Pedagogical University, Tallinn, Estonia: Resting and active evolution. Possible preadaptations in the early evolution of Angiosperms. See also here.

George Poinar and Greg Poinar (2018): The antiquity of floral secretory tissues that provide today’s fragrances. Abstract, Historical Biology. See also:
Schnupperten schon Dinos Blumenduft? Kreidezeitliche Blütenpflanzen könnten bereits Düfte produziert haben In German,

G. Poinar et al. (2016): Fossil species of Boehmerieae Gaudich. (Urticaceae) in Dominican and Mexican amber: A new genus (Ekrixanthera) and two new species with anemophilous pollination by explosive pollen release, and possible lepidopteran herbivory. In PDF, Botany.

D. Pons and D. de Franceschi (2007): Neogene woods from western Peruvian Amazon and palaeoenvironmental interpretation. Bulletin of Geosciences, 82: 343-354.

I. Poole (2000): Fossil angiosperm wood anatomy: its role in the reconstruction of biodiversity and palaeoenvironment. Free access, Botanical journal of the Linnean Society, 134: 361-381.

! J.C. Preston et al. 2022): Plant structure and function: Evolutionary origins and underlying mechanisms. Free access, Plant Physiology.

H. Prier et al. (2004): Exotische Gehölze im KIRCHHEIMER-Arboretum Freiburg. PDF file, in German. LGRB-Informationen, Heft 15 (Landesamt für Geologie, Rohstoffe und Bergbau Baden-Württemberg, Freiburg i. Br.). See also here.

K.M. Pryer et al. (2001): Horsetails and ferns are a monophyletic group and the closest living relatives to seed plants. Abstract, Nature, 409: 618-622.
! See also here (in PDF).

Public Broadcasting Service (PBS): NOVA (a high rated science series on TV). First Flower. Go to: Flowers Modern and Ancient. About Archaefructus liaoningensis, discovered in the fossil beds of Liaoning Province in northeastern China.

! J.-H. Ran et al. (2018): Phylogenomics resolves the deep phylogeny of seed plants and indicates partial convergent or homoplastic evolution between Gnetales and angiosperms. Abstract.

! T. Reichgelt et al. (2018): The relation between global palm distribution and climate. Free access, Scientific Reports, 8:4721, doi:10.1038/s.

D. Ren et al. (2009). A Probable Pollination Mode Before Angiosperms: Eurasian, Long-Proboscid Scorpionflies. In PDF, Science, 326: 840-847. See also here.

D. Ren (1998): Flower-Associated Brachycera Flies as Fossil Evidence for Jurassic Angiosperm Origins. In PDF.
This expired link is still available through the Internet Archive´s Wayback Machine.
See also here.

S.S. Renner (2023): A time tree for the evolution of insect, vertebrate, wind, and water pollination in the angiosperms. Free access, New Phytologist, 240: 464–465.
This article is a Commentary on Stephens et al. (2023), 240: 880–891.

James L. Reveal, Norton-Brown Herbarium, University of Maryland: Advanced Plant Taxonomy. Systems of classification for magnoliophyta, history of systematic botany, approaches to biological classification, taxonomic hierarchy, types of data.

Anita Roth-Nebelsick et al. (2001): Evolution and Function of Leaf Venation Architecture: A Review. PDF file, Annals of Botany 87: 553-566. See also here.

Gar W. Rothwell, Department of Environmental and Plant Biology, Ohio University, Athens: Angiophytes: Using Whole Plant Concepts to Interpret Angiosperm Origins.
Selected Literature.
Links archived by the Internet Archive´s Wayback Machine.

Gar W. Rothwell, Department of Environmental and Plant Biology, Ohio University, Athens, OH: Vascular Plant Morphology. A version archived by Internet Archive Wayback Machine. This course covers the structure, development, reproductive biology and relationships of vascular plants. The course is structured to emphasize the evolutionary changes that led to the diversity of modern tracheophytes. Go to Flowering Plants (PDF file).

D. Royer et al. (2010): Leaf economic traits from fossils support a weedy habit for early angiosperms. Free access, American Journal of Botany, 97: 438-445.

! Dmitry A. Ruban (2012): Mesozoic mass extinctions and angiosperm radiation: does the molecular clock tell something new? In PDF, Geologos, 18: 37-42.

P.J. Rudall and R.M. Bateman (2019): Leaf surface development and the plant fossil record: stomatal patterning in Bennettitales. Abstract, Biological Reviews.
"... Fossil bennettites – even purely vegetative material – can be readily identified by a combination of epidermal features, including distinctive cuticular guard-cell thickenings, lobed abaxial epidermal cells (‘puzzle cells’), transverse orientation of stomata perpendicular to the leaf axis, and a pair of lateral subsidiary cells adjacent to each guard-cell pair (termed paracytic stomata). ..."

P.J. Rudall et al. (2017): Evolution and development of monocot stomata. In PDF, American journal of botany, 104: 1122-1141.

Paula J. Rudall and Richard M. Bateman (2010): Defining the limits of flowers: the challenge of distinguishing between the evolutionary products of simple versus compound strobili. In PDF, Philos. Trans. R. Soc. London, B Biol. Sci., 365: 397-409. See also here (abstract).

! F. Rümpler and G. Theißen (2019): Reconstructing the ancestral flower of extant angiosperms: the ‘war of the whorls’ is heating up. Open access, Journal of Experimental Botany, 70: 2615–2622.
"... we first summarize the results of previous attempts to reconstruct AFEA and contrast them with the more recent, controversial prediction of its structure. We then outline the major arguments made by contrasting parties in the recent debate. Finally, we discuss two key topics, the molecular mechanism of phyllotaxis and the role of gene regulatory networks during flower development and evolution, ..."

A. Salt (2019): When did the first flowers open? Botany One.

A.A. Santos and X. Wang (2022): Pre-Carpels from the Middle Triassic of Spain. Open access, Plants, 11(21), 2833;

H. Sauquet et al. (2022): What is the age of flowering plants? In PDF, Journal of Experimental Botany,
See also here.
Note fig. 1: Hypothetical time tree of the angiosperms.
Note fig. 2. Crown-group angiosperm age estimates obtained in fossil-calibrated molecular dating and palaeontological macroevolutionary modelling studies published over the last 6 years.

! H. Sauquet and S. Magallón (2018): Key questions and challenges in angiosperm macroevolution. In PDF, New Phytologist, 219: 1170–1187. See also here.

! H. Sauquet et al. (2017): The ancestral flower of angiosperms and its early diversification. Free acces, Nature Communications, 8.
Note figure 1: Three-dimensional model of the ancestral flower reconstructed.
"... We reconstruct the ancestral angiosperm flower as bisexual and radially symmetric, with more than two whorls of three separate perianth organs each (undifferentiated tepals), more than two whorls of three separate stamens each, and more than five spirally arranged separate carpels ..."
Also worth checking out:
! Was war die erste Blüte der Erdgeschichte? In German, by Ulf von Rauchhaupt, 2023, Frankfurter Allgemeine Zeitung.

S.R. Schachat 2022): Examining paleobotanical databases: Revisiting trends in angiosperm folivory and unlocking the paleoecological promise of propensity score matching and specification curve analysis. Free access, Front. Ecol. Evol., 10: 951547. doi: 10.3389/fevo.2022.951547.
"... Long-term trends in the fossil record of plants, encompassing their interactions with herbivores and with the environment, are of the utmost relevance for predicting global change
[...] in contrast to modern ecology and unlike various other paleontological disciplines, paleobotany has a limited history of “big data” meta-analyses.
[...] Here I demonstrate the importance of analytical best practices by applying them to a recent meta-analysis of fossil angiosperms. ..."

! H. Schneider et al. (2004): Ferns diversified in the shadow of angiosperms. In PDF, Nature, 428: 553–557. See also here (abstract).
! Note figure 1: Phylogenetic chronograms of ferns (a) and angiosperms (b), and proportional lineages-through-time (LTT) plots for angiosperms and polypods (c).
"... we report divergence time estimates for ferns and angiosperms based on molecular data, with constraints from a reassessment of the fossil record. We show that polypod ferns (>80% of living fern species) diversified in the Cretaceous, after angiosperms ..."

S. Sgorbati et al. (2018): Was Charles Darwin right in his explanation of the ‘abominable mystery’? Free access, Italian Botanist, 5: 25–30.

C. Shi et al. (2022): Fire-prone Rhamnaceae with South African affinities in Cretaceous Myanmar amber. In PDF, Nature Plants, 8: 125–135.
See also here.
"... We report the discovery of two exquisitely preserved fossil flower species, one identical to the inflorescences of the extant crown-eudicot genus Phylica and the other recovered as a sister group to Phylica, both preserved as inclusions together with burned plant remains in Cretaceous amber from northern Myanmar (~99 million years ago) ..."

G. Shi et al. (2021): Mesozoic cupules and the origin of the angiosperm second integument. Abstract, Nature, 594: 223–226. See also here (in PDF).

! D. Silvestro et al. (2020): Fossil data support a pre-Cretaceous origin of flowering plants. In PDF, Nature Ecology & Evolution. See also here.
"... Yet, our results indicate that an early, pre-Cretaceous origin of angiosperms is supported not only by molecular phylogenetic hypotheses but also by an analysis of the fossil record ..."

! Stephen A. Smith et al. (2010): An uncorrelated relaxed-clock analysis suggests an earlier origin for flowering plants. PDF file, PNAS, 107: 5897-5902. See also here, or there.

! S.A. Smith and M.J. Donoghue (2008): History in Flowering Plants Rates of Molecular Evolution Are Linked to Life. In PDF, Science, 322. See also here (abstract).

D.D. Sokoloff et al. (2019): Supposed Jurassic angiosperms lack pentamery, an important angiosperm-specific feature. Open access, New Phytologist.

D. Soltis et al. (2017): Phylogeny and Evolution of the Angiosperms. Book announcement. See also here (Google books). Worth checking out:
! Relationships of Angiosperms to Other Seed Plants. In PDF.

P.S. Soltis et al. (2019): Darwin review: angiosperm phylogeny and evolutionary radiations. In PDF, Proc. R. Soc. B, 286: 20190099. See also here.

! D.E. Soltis et al. (2009): Polyploidy and angiosperm diversification. Free access, Am. J. Bot., 96: 336-348.

P. Soltis, (a resource of the American Institute of Biological Sciences): Flowering Plants: Keys to Earth´s Evolution and Human Well-Being. A version archived by Internet Archive Wayback Machine.

P.S. Soltis and D.E. Soltis (2004): The origin and diversification of angiosperms. Free access, American Journal of Botany, 91: 1614-1626.

Space Daily: Paleobotanist identifies what could be the mythical "first flower". (August 18, 2015).

! Doug Soltis, Amber Tilley and Hongshan Wang, Florida Museum of Natural History (FLMNH), University of Florida: Deep Time. A comprehensive phylogenetic tree of living and fossil angiosperms. Deep Time explore the ways in which angiosperm fossils can be appropriately integrated into the phylogenetic framework for extant taxa, with the ultimate goal of forming a comprehensive phylogenetic tree of living and fossil angiosperms. This includes the evaluation and prioritization of the fossil record, the critical appraisal of the age of fossils, the construction of a morphological data matrix for fossils and extant angiosperms, the integration of fossils into the angiosperm tree and the calibration of divergence times.

! R.E. Stephens et al. (2023): Insect pollination for most of angiosperm evolutionary history. Open access, New Phytologist, 240: 880–891.
"... Most contemporary angiosperms (flowering plants) are insect pollinated, but pollination by wind, water or vertebrates occurs in many lineages.
[...] We use a robust, dated phylogeny and species-level sampling across all angiosperm families to model the evolution of pollination modes
[...] Angiosperms were ancestrally insect pollinated, and insects have pollinated angiosperms for c. 86% of angiosperm evolutionary history ..."

! P.F. Stevens and Hilary Davis, Missouri Botanical Garden, St. Louis: Angiosperm Phylogeny Website. The focus of this site is on angiosperms and emphasis is placed on plant families. You can also navigate from the Orders- or the Families-website. Go to:

! P.F. Stevens and Hilary Davis, Missouri Botanical Garden, St. Louis: Angiosperm Phylogeny Website. The focus of this website is about classification on angiosperm plant families. Sets of evolutionary trees grouping families and orders and showing details of the arrangement. Don´t miss the complete synonymy of family name directory and some literature references. Excellent!

Ruth A. Stockey, Department of Biological Sciences, University of Alberta, Edmonton: Paleobotany of Angiosperm Origins. Go to: Course Outline. Chiefly bibliographies and weblinks.
These expired links are now available through the Internet Archive´s Wayback Machine.

G. Sun et al. (1998): In Search of the First Flower: A Jurassic Angiosperm, Archaefructus, from Northeast China. In PDF, Science, 282: 1692-1695.
Images of Archaefructus liaoningensis.

Syllabus of Plant Families, A. Engler´s Syllabus der Pflanzenfamilien (13th edition by Wolfgang Frey):
4 Pinopsida (Gymnosperms), Magnoliopsida (Angiosperms) p.p.: Subclass Magnoliidae [Amborellanae to Magnolianae, Lilianae p.p. (Acorales to Asparagales)].

Ralph E. Taggart, Department of Botany and Plant Pathology/Department of Geological Sciences at Michigan State University, East Lansing:
! BOT335 Lecture Schedule. Some interesting chapters in terms of palaeobotany, e.g.
The First Vascular Land Plants;
Carboniferous Forests;
Arborescent Lycopods;
Psaronius: a Carboniferous tree-fern;
Carboniferous Horsetails;
Carboniferous Seed Ferns;
The Evolution of Conifers;
Cycadophytes, the True Cycads;
Mesozoic Cycadeoids;
North American Redwoods, Past and Present.
These expired links are available through the Internet Archive´s Wayback Machine.

D.W. Taylor and H. Li (2018): Paleobotany: Did flowering plants exist in the Jurassic period? eLife, 7: e43421.
"... we infer that Nanjinganthus shows substantial similarity to predicted models of ancestral characters and Early Cretaceous angiosperms, so the evidence suggests that it is a Jurassic flowering plant. ..."

David Winship Taylor et al. (2006): Biogeochemical evidence for the presence of the angiosperm molecular fossil oleanane in Paleozoic and Mesozoic non-angiospermous fossils. Abstract, Paleobiology, 32: 179-190.

! David W. Taylor and Leo J. Hickey (1996): Flowering Plant Origin, Evolution & Phylogeny. Google books (some pages omitted); American Institute of Biological Sciences (Springer), 404 pages.

! E.L. Taylor and T.N. Taylor (2009): Seed ferns from the late Paleozoic and Mesozoic: Any angiosperm ancestors lurking there? Open access, American Journal of Botany, 96: 237-251.
! "... In our opinion, it will be more productive to attempt to solve Darwin’s mystery if there were greater attention directed at mining the rock record in the hope of discovering more informative and new specimens, than to continue to construct new phylogenies using the same, often ambiguous characters. ..."
Worth checking out: Glossopterid vegetative and reproductive organs:
Note fig. 2: Suggested reconstruction of Ottokaria zeilleri.
Fig. 10: Suggested reconstruction of a Glossopteris megasporophyll with seeds attached to adaxial surface.
12: Diagrammatic reconstruction of Denkania indica.
Reproductive organs of Caytoniales and Corystospermales:
15. Suggested reconstruction of Caytonia cupule showing attachment of seeds and “stigmatic lip”.
16. Reconstruction of Caytonanthus arberi.
19. Suggested reconstruction of Umkomasia asiatica.
21. Diagrammatic reconstruction of Umkomasia uniramia.
Reproductive organs of Corystosperms and Petriellales:
25. Suggested reconstruction of Pilophorosperma geminatum.
28. Suggested reconstruction of Pteruchus fremouwensis.
30. Suggested reconstruction of Petriellaea triangulata.
32. Diagrammatic cutaway of Petriellaea triangulata cupule.
Reproductive organs of peltasperms:
34. Suggested reconstruction of Autunia conferta ovuliferous organ.
36. Suggested reconstruction of two Autunia conferta megasporophylls.
37. Suggested reconstruction of Peltaspermum rotula megasporophyll showing several ovules.
39. Suggested reconstruction of Peltaspermum thomasii axis bearing numerous megasporophylls.
40. Suggested reconstruction of Peltaspermopsis polyspermis.
41. Suggested reconstruction of Lepidopteris frond with pollen organs of the Antevsia-type at the tip.
42. Suggested reconstruction of Antevsia zeilleri pollen organ showing pinnate axis bearing clusters of pollen sacs.

L.B. Thien et al. (2000): New Perspectives on the Pollination Biology of Basal Angiosperms. Abstract, International Journal of Plant Sciences, 161.

H. Hamshaw Thomas (1936): Palaeobotany and the origin of the angiosperms. First page, The Botanical Review, 2: 397-418. See also here.

J.B. Thompson and S. Ramírez-Barahona (2023): No evidence for angiosperm mass extinction at the Cretaceous–Paleogene (K–Pg) boundary. In PDF, bioRxiv.

Greg Thorn, Department of Biology, University of Western Ontario, Canada:
! Evolution of Plants . Lecture notes, e.g.:
Evolution of the Angiosperms. Powerpoint presentation.

Greg Thorn, Department of Biology, University of Western Ontario: Evolution of Plants (Powerpoint presentations). Navigate from here with information from the Syllabus. See e.g. Lecture 16: Evolution of Plants. The evolution of early angiosperms.

! B.H. Tiffney and S.R. Manchester (2001): The Use of Geological and Paleontological Evidence in Evaluating Plant Phylogeographic Hypotheses in the Northern Hemisphere Tertiary. Abstract, International Journal of Plant Sciences, 162. See also here (in PDF).

! Amber Tilley and Hongshan Wang, Florida Museum of Natural History, University of Florida: Deep Time. Projects about the origin of angiosperms.

Tree of Life Web Project: Angiosperms (by Pam Soltis, Doug Soltis, and Christine Edwards).

G.R. Upchurch Jr. (1984): Cuticle evolution in Early Cretaceous angiosperms from the Potomac Group of Virginia and Maryland. PDF file, Annals of the Missouri Botanical Garden. A version archived by Internet Archive Wayback Machine.

! C.J. van der Kooi and J. Ollerton (2020): The origins of flowering plants and pollinators. Free access, Science, 368: 1306-1308.
See also here (in PDF).

B. Vento et al. (2023): Phylogenetic relationships in Nothofagus: The role of Antarctic fossil leaves. In PDF, Acta Palaeontologica Polonica, 68.

S. Wang et al. (2021): Cretaceous fire-resistant angiosperms. In PDF, preprint, DOI:
See also here.
"... both preserved as inclusions in Cretaceous amber from northern Myanmar (~99 Ma). These specialized flower structures, named Phylica piloburmensis sp. nov. and Eophylica priscastellata gen. et sp. nov., were adapted to surviving frequent wildfires, providing the earliest evidence of fire-resistance in angiosperms. ..."

H. Wang et al. (2013): Fruits, seeds, and flowers from the Warman clay pit (middle Eocene Claiborne Group), western Tennessee, USA. In PDF, Palaeontologia Electronica. See also here.

X. Wang (2023): Origin of Angiosperms: Problems, Challenges, and Solutions. Free access, Life, 13.

X. Wang: An Era of Errors. About Archaeanthus (in PDF). See also here.

! X. Wang (2017): A Biased, Misleading Review on Early Angiosperms. In PDF, Natural Science, 9: 399-405.
Please note: P.S. Herendeen et al. (2017): Palaeobotanical redux: revisiting the age of the angiosperms. In PDF, Nature Plants 3. See also here.

X. Wang (2010): Suggested Angiosperm Ancestors. In PDF, Chapter 2, The Dawn Angiosperms. Lecture Notes in Earth Sciences Volume 121: 5-16. See also here (Springer) and there Google books.

X. Wang et al. (2007): Schmeissneria: A missing link to angiosperms? Open access, BMC Evolutionary Biology, 7. See also here.

X.A. Wang (2022): A Novel Early Cretaceous Flower and Its Implications on Flower Derivation. Free access, Biology, 11.

! L. Watson Albany, Australia, and M. J. Dallwitz CSIRO Entomology, Canberra, Australia (page hosted by DELTA): The Families of Flowering Plants. Descriptions, illustrations, identification, and information retrieval. Version: 25th November 2009. The Intkey software is required.
Also worth to check out: Static information. Character list, implicit attributes, notes on the APG classification, etc.
Snapshot taken by the Internet Archive´s Wayback Machine.

G.D.A. Werner et al. (2014): A single evolutionary innovation drives the deep evolution of symbiotic N2-fixation in angiosperms. Open access, Nature Communications, 5: 4087.

Biology Department, Western Washington University, Bellingham, Washington:
! Flowering vascular plants. Powerpoint presentation. See also here, or there.

E.A. Wheeler and S.R. Manchester (2007): Review of the wood anatomy of extant Ulmaceae as context for new reports of late Eocene Ulmus woods. PDF file, Bulletin of Geosciences, 82: 329-342.

! E.A. Wheeler and P. Baas (2022): Wood anatomy of modern and fossil Fagales in relation to phylogenetic hypotheses, familial classification, and patterns of character evolution. Free access, International Journal of Plant Sciences, 183.

! E.A. Wheeler and P. Baas (1991): A Survey of the Fossil Record for Dicotiledonous Wood and its Significance for Evolutionary and Ecological Wood Anatomy. Free access, IAWA Bulletin n.s., 12: 275-332.
Note figure 1: Major ecophyletic trends of vessel element specialisation.

Niklas Wikström et al. (2001): Evolution of the angiosperms: calibrating the family tree. PDF file, Proc. R. Soc. Lond., B, 268: 2211-2220.

! Wikipedia, the free-content encyclopedia: Spermatophyte. Go to: Flowering Plant, and Magnoliopsida.

P. Wilf et al. (2023): The end-Cretaceous plant extinction: Heterogeneity, ecosystem transformation, and insights for the future. Open access, Cambridge Prisms: Extinction, 1, e14, 1–10.

P. Wilf (2008): Fossil angiosperm leaves: paleobotany´s difficult children prove themselves. PDF file, Paleontological Society Papers, 14: 319-333.

Kathy Willis and Jennifer McElwain: The Evolution of Plants. Oxford University Press, Second Edition. Don't miss the
Companion Website
and some samples in Google books.
Note chapter 1: The evolutionary record and methods of reconstruction (in PDF).

Kathy Willis, School of Geography and the Environment, University of Oxford, & Jenny McElwain, Field Museum of Natural History, Chicago (Oxford University Press): The Evolution of Plants. Book announcement. Go to: Chapter 06, Flowering plant origins (PDF file).
Snapshots provided by the Internet Archive´s Wayback Machine.

! Hugh D. Wilson, Department of Biology Herbarium (TAMU), Texas A&M University: Flowering Plant Gateway. This project involves the development of computer programs that allow automated, machine-generated HTML page production for each Subclass/Superorder of the Flowering Plants as structured by the Cronquist, Takhtajan, and Thorne Systems of classification.
Snapshot provided by the Internet Archive´s Wayback Machine.

! Hugh D. Wilson, Department of Biology Herbarium (TAMU), Texas A&M University: Flowering Plant Gateway.
This project involves the development of computer programs that allow automated, machine-generated HTML page production for each Subclass/Superorder of the Flowering Plants as structured by the Cronquist, Takhtajan, and Thorne Systems of classification.

! S.L. Wing and L.D. Boucher (1998): Ecological aspects of the Cretaceous flowering plant radiation. In PDF, Annu. Rev. Earth Planet. Sci. 1998 26: 379-421.

! A.E. Zanne et al. (2014): Three keys to the radiation of angiosperms into freezing environments. In PDF, Nature. Provided by the Internet Archive´s Wayback Machine.

L. Zhang et al. (2020): The water lily genome and the early evolution of flowering plants. Open access, Nature, 577: 79–84. Worth checking out:
Fig. 1d: Summary phylogeny and timescale of 115 plant species. Blue bars at nodes represent 95% credibility intervals of the estimated dates.

X. Zhang et al. (2017): How the ovules get enclosed in magnoliaceous carpels. PLoS ONE, 12: e0174955.

! A.R. Zuntini et al. (2024): Phylogenomics and the rise of the angiosperms. Free access, Nature.
"... we build the tree of life for almost 8,000 (about 60%) angiosperm genera
[...] Scaling this tree to time using 200 fossils, we discovered that early angiosperm evolution was characterized by high gene tree conflict and explosive diversification, giving rise to more than 80% of extant angiosperm orders. Steady diversification ensued through the remaining Mesozoic Era until rates resurged in the Cenozoic Era, concurrent with decreasing global temperatures
[...] our extensive sampling combined with advanced phylogenomic methods shows the deep history and full complexity in the evolution of a megadiverse clade ..."

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